The phylogeny of the Noctuidae...

Systematic Entomology(1996)21, 219–251

The phylogeny of the Noctuidae (Lepidoptera)

W. S P E I D E L , H . FAN G E R and C . M . N A U M A N N Zoologisches Forschungsinstitut und

Museum Alexander Koenig, Bonn, Germany

Abstract. A new phylogenetic system of the Noctuidae is proposed. The systemshould, however, be regarded as provisional because it is primarily based on only twocharacter complexes: the male genitalia and the tympanal region. The presence ofpreabdominal brush-organs and the length of tibial spurs in the adult male, thepresence of a ventral cervical gland and the lack of the SV 2 setae on the firstabdominal segment of the larvae are considered. The results confirm the monophylyof the Noctuidae. The Herminiinae are considered to represent the sister-group of allother Noctuidae. Remaining noctuid subfamilies are grouped in a series of clades forwhich the corresponding autapomorphies are given. Some formerly acceptedsubfamilies, such as the Acontiinae, prove to be polyphyletic. Accordingly, thesesubfamilies are subdivided into separate lineages. A sister-group relationship betweenthe Noctuidae and the Aganaidae is supported. These two families may be unitedwith the Arctiidae to form a still broader clade.

Introduction superficial resemblance and vaguely defined characters, ratherthan on rigorous application of cladistic principles.

In an attempt to progress beyond the age of traditionalSeven moth families, the Oenosandridae, Doidae,noctuid taxonomy we started a morphological investigation,Notodontidae, Lymantriidae, Arctiidae, Aganaidae andbased on a set of characters, which we considered particularlyNoctuidae, are currently recognized in the large superfamilyuseful in elucidating the basic relationships of the noctuidNoctuoidea (Miller, 1991), comprising some 40 000 describedsubfamilies.species (Heppner, 1991). Monophyly of the Noctuoidea is

The tympanal region was chosen because it was demonstratedbased on the presence of a metathoracic tympanal organ (Miller,by Eggers (1919) to be structurally heterogeneous within the1991). With almost 25 000 included species, the NoctuidaeNoctuidae. Richards (1932) proposed a noctuid classificationform the largest family within the superfamily, and within thebased on a comparative study of the tympanal region.order Lepidoptera as a whole (Heppner, 1991). Nevertheless,Unfortunately, Richard’s system relies exclusively on intuitiveour understanding of the phylogenetic affinities of the Noctuidaecharacter interpretation; characters or character states to delimitto other noctuoid families, as well as interrelations among thesubfamily groups were not provided.various noctuid subfamilies, is still in its infancy.

The male genitalia and their associated muscles wereHolloway (1988) noted that the Aganaidae and Noctuidaeconsidered because they are thought to evolve quicklyshare a similar number of thoracic subventral setae in the(Eberhard, 1985), and may thus provide considerablelarvae, and both have an orbicular stigma in the forewing cell.information. Tikhomirov (1979a,b,c), in a series of valuableOn these grounds he relegated the Aganaidae to a subfamilypapers, has demonstrated that some noctuid subfamilies shareof the Noctuidae. However, there is homoplasy in thethe unique presence of an additional (novel) male genitalicdistribution of the subventral setae among the families undermuscle.consideration. We prefer to retain family rank for the Aganaidae,

The males of most ‘higher’ Noctuidae are characterized bybecause this corresponds better with the historical developmentthe presence of a paired brush-organ, situated on the secondof noctuoid classification, and because the sister-group relation-abdominal segment, that is presumably responsible for theship is still weakly supported.dissemination of male sexual pheromones (reviewed in BirchKitching (1984) published an historical review of noctuidet al., 1990). We investigated the distribution of this charactersubfamily relationships. His synopsis shows that current ideasin order to find the point at which it evolved within the family.on the higher classification of the Noctuidae are based on

During our studies we suspected that the spurs of noctuidspecies might be relatively long. We therefore compared spurlength of the Noctuidae with that of outgroups.Correspondence: Dr W. Speidel, Zoologisches Forschungsinstitut

The presence of a ventral cervical gland in noctuid larvaeund Museum Alexander Koenig, Adenauerallee 160, D-53113 Bonn,Germany. has been known for quite a long time (e.g. Gardner, 1941;

© 1996 Blackwell Science Ltd 219

220 W. Speidel, H. Fa¨nger and C. M. Naumann

Peterson, 1962). Curiously, the precise distribution of this Character system 1: the tympanal regioncharacter within Noctuoidea has never been investigated. Wetherefore decided to include this character in our analysis. General description

The absence of seta SV 2 on the first abdominal segment ofthe larva has been considered to represent an apomorphic The term tympanal region is used here in a rather broadcharacter for the higher Noctuidae (Poole, 1995). We also sense. It refers to the area around the waist-like cleft separatingincluded this important character. the thorax and abdomen, commonly known in the Noctuoidea

as the tympanal cavity (Figs 2 and 3). Thus, the tympanalregion includes the tympanal organ proper as well asfunctionally related structural modifications of the metathoraxMaterial and methodsand first abdominal segment. Sexual dimorphism in tympanalmorphology is probably restricted to species with winglessFor the study of cuticular structures, the specimens werefemales, e.g. the lymantriid genusOrgyia (Eggers, 1919). Thecleared in hot aqueous KOH (10%) for about 10 min. In somesex of the specimens investigated is therefore not noted.cases the specimens were later stained in an aqueous solution of

We focus on external structures of the tympanal region. AnEvan’s Blue in order to clearly visualize delicate membraneousoverview of structures and morphological terms is presentedstructures.in Fig. 2, showing the metathorax and the first two abdominalSpecimens to be studied by scanning electron microscopysegments of a higher noctuid species,Agrotis exclamationis(SEM) were either macerated in KOH, or fixed in an aqueous(Noctuinae). The following structures are of particular interest:solution of picric acid. In both cases they were dehydrated

through a graded series of ethanol before being transferred tohexamethyldisilazan. Specimens were then air-dried and

1 Hood structures of the A1 pleuronsputter-coated with gold prior to SEM examination.

For the investigation of genitalic muscles, male moths wereThe presence of a metathoracic tympanum has influencedstored in a saturated aqueous solution of picric acid until

the structure of the pleural region of the first abdominaldissection. Penetration of the aqueous solution into thesegment (A1). The anterior parts of A1, forming the posteriorspecimens was greatly facilitated by first soaking them brieflywall of the tympanal cavity, are often prominently expanded.in 70% ethanol. Prolonged storage in picric acid stains theThere are three principal configurations. A prespiracular hood,muscles a bright yellow colour, while they simultaneouslyi.e. a pleural evagination situated anterior to the A1 spiracle,remain soft and flexible. During dissection, which was carriedis present in the Oenosandridae, Lymantriidae, Arctiidae andout in 70% ethanol, specimens were occasionally dipped in ain a strongly reduced form in the Aganaidae (Fig. 4). Thediluted alcoholic solution of Evan’s Blue in order to obtainHerminiinae are the only noctuids possessing this configurationenhanced contrast.(Fig. 5). A postspiracular hood originating immediately behindWe obtained fixed male specimens from the noctuoid familiesthe A1 spiracle is typical for the remaining Noctuidae (Figs 2,Oenosandridae, Notodontidae, Lymantriidae, Arctiidae,6, 7). Reduction of the hood to simple dorsal remnants occursAganaidae and from most subfamilies of the Noctuidaein the Nolinae (Fig. 8). Finally, a perispiracular cup on the A1(Table 1a). Unfortunately, we were unable to obtain fixed malepleura is typical for all Notodontidae (Fig. 9). In some casesspecimens of the Doidae, a small family confined to thethe cup has shifted anterad to such an extent that it bearsAmericas.superficial resemblance to a postspiracular hood, and may evenA list of the species examined for the other characters isbe mistaken for that structure.given on Table 1(b–d).

In addition, the pleural region posterior to the A1 hood is,at least in the Noctuidae, provided with a lateral sclerotization(Figs 5–7). This lateral A1 sclerotization should not be confused

Results with the hood itself. This sclerotization may be anautapomorphy of the Noctuidae1 Aganaidae (Minet, 1986;

At the outset, the phylogenetic system we propose as a resultcharacter a), but has not been studied here in detail.of our morphological investigations will be briefly introduced.Family relationships between the Arctiidae, Aganaidae and

2 The alulaNoctuidae, and the principal noctuid subfamily arrangements,are depicted in Fig. 1. For each taxon the approximate numberof species has been added in parentheses, mainly following The alula, the posterobasal lobe of the wing, forms a thin

membraneous flap covering the dorsal part of the tympanalHeppner (1991). Information concerning the monophyly andcomposition of individual noctuid subfamilies is presented later region. Its tubular posterior margin (axillary cord) is continued

into the scutellar arm, connecting the hindwing with thein the discussion. The cladogram is accompanied by a list ofthe presumed autapomorphies for each of the ten clades metanotum (Fig. 14). Generally, the insect metanotum houses

a pulsatile wing-heart that is responsible for suckingindicated (Table 2). In the following pages the apomorphies inTable 2 are detailed through successive treatment of the five haemolymph out of the hindwings along the scutellar arm (e.g.

Krenn & Pass, 1994, 1995). Because of its large size the alulacharacter systems examined.

© 1996 Blackwell Science Ltd,Systematic Entomology, 21, 219–251

Phylogeny of Noctuidae 221

Table 1 List of families and species investigated.

(a) For the survey of the muscles in the male genitalia*Oenosandridae

Oenosandra boisduvaliiNewman, 1856Notodontidae

Cerura vinula(Linnaeus, 1758)Phalera bucephala(Linnaeus, 1758)Thaumatopoea solitaria(Freyer, 1838)Traumatocampa pityocampa([Denis & Schiffermuller], 1775)

LymantriidaeCalliteara pudibunda(Linnaeus, 1758)Lymantria dispar(Linnaeus, 1758)

ArctiidaeEilema complana(Linnaeus, 1758)Euplagia quadripunctaria(Poda, 1761)

AganaidaeAsota caricae(Fabricius, 1775)

NoctuidaeHerminia grisealis([Denis & Schiffermuller], 1775) (Herminiinae)Hydrillodes moloalis(Walker, 1859) (Herminiinae?)Rivula sericealis(Scopoli, 1763) (Rivulinae)Hypena proboscidalis(Linnaeus, 1758) (Hypeninae)Penicillaria simplex(Walker, 1865) (Euteliinae)Stictoptera cucullioidesGuenee, 1852 (Stictopterinae)Catocala fraxini(Linnaeus, 1758) (Catocalinae)Oxyodes scrobiculata(Fabricius, 1775) (Catocalinae?)Panthea coenobita(Esper, 1785) (Pantheinae)Colocasia coryli(Linnaeus, 1758) (Pantheinae)Diloba caeruleocephala(Linnaeus, 1758) (Pantheinae)Camptoloma interiorata(Walker, 1865) (Camptolominae)Maurilia iconica (Walker, 1858) (Chloephorinae)Pseudoips prasinana(Linnaeus, 1758) [5 fagana(Fabricius, 1781)] (Chloephorinae)Blenina lucretia(Dalman, 1823) [5 accipiensWalker, [1858]] (Sarrothripinae)Blenina quinariaMoore, 1882 (Sarrothripinae)Meganola albula([Denis & Schiffermuller], 1775) (Nolinae)Meganola strigula([Denis & Schiffermuller], 1775) (Nolinae)Abrostola triplasia(Linnaeus, 1758) [5 trigemina (Werneburg, 1864)] (Plusiinae)Autographa gamma(Linnaeus, 1758) (Plusiinae)Acronicta rumicis(Linnaeus, 1758) (Acronictinae)Cryphia algae(Fabricius, 1775) (Bryophilinae)Sarbanissasp. (Agaristinae)Heliothis virescens(Fabricius, 1777) (Heliothinae)Noctua pronubaLinnaeus, 1758 (Noctuinae)Hecatera bicolorata(Hufnagel, 1766) (Hadeninae)

(b) For the survey of the presence of a ventral cervical gland in the larvaeNotodontidae

Phalera bucephala(Linnaeus, 1758)Pheosia gnoma(Fabricius, 1776)Stauropus fagi(Linnaeus, 1758)

LymantriidaeLymantria dispar(Linnaeus, 1758)Orgyia sp.

ArctiidaeArctia caja (Linnaeus, 1758)Divarctia diva (Staudinger, 1887)Eilema caniola(Hubner, [1808])Phragmatobia fuliginosa(Linnaeus, 1758)Spilosoma lubricipeda(Linnaeus, 1758)

AganaidaeAsota planaWalker, 1854



Table 1 Continued.

NoctuidaeHerminia tarsicrinalis(Knoch, 1782) (Herminiinae)Pechipogo strigilataLinnaeus, 1758 (Herminiinae)Rivula sericealis(Scopoli, 1763) (Rivulinae)Schrankia taenialis(Hubner, [1809]) (Hypenodinae)Schrankia costaestrigalis(Stephens, 1834) (Hypenodinae)Hypena proboscidalis(Linnaeus, 1758) (Hypeninae)Hypena rostralis(Linnaeus, 1758) (Hypeninae)Eutelia adulatrix(Hubner, [1813]) (Euteliinae)Calyptra thalictri (Borkhausen, 1790) (Catocalinae)Catocala promissa([Denis & Schiffermuller], 1775) (Catocalinae)Dysgonia algira(Linnaeus, 1767) (Catocalinae)Colocasia coryli(Linnaeus, 1758) (Pantheinae)Diloba caeruleocephala(Linnaeus, 1758) (Pantheinae)Pseudoips prasinana(Linnaeus, 1758) (Chloephorinae)Nycteola revayana(Scopoli, 1772) (Sarrothripinae)Eligma narcissus(Cramer, 1776) (Sarrothripinae)Autographa gamma(Linnaeus, 1758) (Plusiinae)Protodeltote pygarga(Hufnagel, 1766) (Eustrotiinae)Eublemma minutata(Fabricius, 1794) (Acontiinae?)Phlogophora meticulosa(Linnaeus, 1758) (Amphipyrinae)Omphaloscelis lunosa(Haworth, 1809) (Amphipyrinae)Discestra trifolii (Hufnagel, 1766) (Hadeninae)

(c) For the survey of the tympanal regionOenosandridae

Oenosandra boisduvaliiNewman, 1856Notodontidae

Cerura vinula(Linnaeus, 1758)Drymonia obliterata(Esper, 1785)Leucodonta bicoloria([Denis & Schiffermuller], 1775)Phalera bucephala(Linnaeus, 1758)Pheosia gnoma(Fabricius, 1776)Thaumatopoea solitaria(Freyer, 1838)

LymantriidaeCalliteara pudibunda(Linnaeus, 1758)Lymantria dispar(Linnaeus, 1758)

ArctiidaeCreatonotus transiens(Walker, 1855)Dysauxes ancilla(Linnaeus, 1767)Eilema complana(Linnaeus, 1758)Euplagia quadripunctaria(Poda, 1761)Miltochrista miniata(Forster, 1771)Tyria jacobeae(Linnaeus, 1758)

AganaidaeAsota caricae(Fabricius, 1775)Asota egens(Walker, 1854)Digama hearseyanaMoore, [1860]

NoctuidaeAdrapsa ablualis(Walker, [1859]) (Herminiinae)Herminia grisealis([Denis & Schiffermuller], 1775) (Herminiinae)Herminia ryukyuensisOwada, 1982 (Herminiinae)Herminia tarsicrinalis(Knoch, 1782) (Herminiinae)Hydrillodes moloalis(Walker, 1859) (Herminiinae?)Nodaria tristis (Butler, 1879) (Herminiinae)Simplicia caeneusalis(Walker, [1859]) (Herminiinae)Trisateles emortualis([Denis & Schiffermuller], 1775) (Herminiinae)Zanclognatha yakushimalisSugi, 1959 (Herminiinae)Rivula sericealis(Scopoli, 1763) (Rivulinae)Anoratha costalisMoore, 1867 (Hypeninae)Hypena crassalis(Fabricius, 1787) (Hypeninae)



Table 1 Continued.

Hypena proboscidalis(Linnaeus, 1758) (Hypeninae)Eutelia adulatrix(Hubner, [1813]) (Euteliinae)Penicillaria simplex(Walker, 1865) (Euteliinae)Stictoptera cucullioidesGuenee, 1852 (Stictopterinae)Blasticorhinussp. (Catocalinae)Catephia alchymista([Denis & Schiffermuller], 1775) (Catocalinae)Catocala fraxini(Linnaeus, 1758) (Catocalinae)Lygephila craccae([Denis & Schiffermuller], 1775) (Catocalinae)Lygephila pastinum(Treitschke, 1826) (Catocalinae)Oxyodes scrobiculata(Fabricius, 1775) (Catocalinae?)Scoliopteryx libatrix(Linnaeus, 1758) (Catocalinae)Ulotrichopus macula(Hampson, 1891) (Catocalinae)Panthea coenobita(Esper, 1785) (Pantheinae)Colocasia coryli(Linnaeus, 1758) (Pantheinae)Diloba caeruleocephala(Linnaeus, 1758) (Pantheinae)Euromoia subpulchra(Alpheraky, 1897) (Pantheinae?)Raphia fraterGrote, 1864 (Pantheinae)Moma alpium(Osbeck, 1778) (Pantheinae)Trichosea ludifica(Linnaeus, 1758) (Pantheinae)Camptoloma interiorata(Walker, 1865) (Camptolominae)Maurilia iconica (Walker, 1858) (Chloephorinae)Bena bicolorana(Fuessli, 1775) (Chloephorinae)Pseudoips prasinana(Linnaeus, 1758) [5 fagana(Fabricius, 1781)] (Chloephorinae)Eligma narcissus(Cramer, 1776) (Sarrothripinae)Nycteola revayana(Scopoli, 1772) (Sarrothripinae)Risoba prominensMoore, 1881 (Sarrothripinae)Meganola albula([Denis & Schiffermuller], 1775) (Nolinae)Meganola strigula([Denis & Schiffermuller], 1775) (Nolinae)Nola cucullatella(Linnaeus, 1758) (Nolinae)Armada panaceorum(Menetries, 1849) (Eustrotiinae?)Protodeltote pygarga(Hufnagel, 1766) (Eustrotiinae)Abrostola triplasia(Linnaeus, 1758) [5 trigemina (Werneburg, 1864)] (Plusiinae)Autographa gamma(Linnaeus, 1758) (Plusiinae)Diachrysia chrysitis(Linnaeus, 1758) (Plusiinae)Acontia lucida(Hufnagel, 1766) (Acontiinae)Emmelia trabealis(Scopoli, 1763) (Acontiinae)Aedia funesta(Esper, 1786) (Aediinae)Tyta luctuosa([Denis & Schiffermuller], 1775) (Tytinae)Aegle koekeritziana(Hubner, 1799) (Stiriinae)Panemeria tenebrata(Scopoli, 1763) (Stiriinae?)Acronicta leporina(Linnaeus, 1758) (Acronictinae)Acronicta rumicis(Linnaeus, 1758) (Acronictinae)Nacna malachitis(Oberthur, 1880) [5 splendens(Moore, 1888)] (Acronictinae)Cryphia algae(Fabricius, 1775) (Bryophilinae)Sinocharis korbaePungeler, 1912 (Sinocharinae)Pseudeustrotia candidula([Denis & Schiffermuller], 1775) (Pseudeustrotiinae)Sarbanissasp. (Agaristinae)Heliothis virescens(Fabricius, 1777) (Heliothinae)Agrotis exclamationis(Linnaeus, 1758) (Noctuinae)Agrotis segetum([Denis & Schiffermuller], 1775) (Noctuinae)Noctua janthina([Denis & Schiffermuller], 1775) (Noctuinae)Cucullia umbratica(Linnaeus, 1758) (Cuculliinae)Shargacucullia scrophulariae([Denis & Schiffermuller], 1775) (Cuculliinae)Hecatera bicolorata(Hufnagel, 1766) (Hadeninae)Mythimna impura(Hubner, [1808]) (Hadeninae)Agrochola helvola(Linnaeus, 1758) (Amphipyrinae)Amphipyra pyramidea(Linnaeus, 1758 (Amphipyrinae)Conistra erythrocephala([Denis & Schiffermuller], 1775) (Amphipyrinae)Cosmia trapezina(Linnaeus, 1758) (Amphipyrinae)Hoplodrina ambigua([Denis & Schiffermuller], 1775) (Amphipyrinae)Mesoligia furuncula([Denis & Schiffermuller], 1775) (Amphipyrinae)



Table 1 Continued.

Omphaloscelis lunosa(Haworth, 1809) (Amphipyrinae)Paradrina clavipalpis(Scopoli, 1763) (Amphipyrinae)Polymixis argillaceago(Hubner, [1822]) (Amphipyrinae)

(d) For the survey of the relative spur lengthGeometridae

Operophthera brumata(Linnaeus, 1758)Hesperiidae

Thymelicus sylvestris(Poda, 1761)Pieridae

Pieris napi (Linnaeus, 1758)Lasiocampidae

Eriogaster lanestris(Linnaeus, 1758)Malacosoma castrense(Linnaeus, 1758)

SaturniidaeAglia tau (Linnaeus, 1758)Saturnia pyri([Denis & Schiffermuller], 1775)

SphingidaeMimas tiliae (Linnaeus, 1758)Deilephila porcellus(Linnaeus, 1758)

OenosandridaeOenosandra boisduvaliiNewman, 1856

NotodontidaeClostera anachoreta([Denis & Schiffermuller], 1775)Thaumatopoea solitaria(Freyer, 1838)

LymantriidaeArctornis l-nigrum(Muller, 1764)Dasychira dudgeoniSwinhoe, 1907Lymantria dispar(Linnaeus, 1758)

ArctiidaeLithosia quadra(Linnaeus, 1758)Euplagia quadripunctaria(Poda, 1761)Phragmatobia fuliginosa(Linnaeus, 1758)Spilosoma lubricipedum(Linnaeus, 1758)Amata phegea(Linnaeus, 1758)

AganaidaeAgape chloropyga(Walker, 1854)Asota caricae(Fabricius, 1775)Asota heliconia(Linnaeus, 1758)Digama hearseyanaMoore, [1860]

NoctuidaeZanclognatha tarsipennalis(Treitschke, 1835) (Herminiinae)Rivula sericealis(Scopoli, 1763) (Rivulinae)Hypena crassalis(Fabricius, 1787) (HypeninaeEutelia adulatrix(Hubner, [1813]) (Euteliinae)Stictoptera semialba(Walker, 1864) (Stictopterinae)Catocala nupta(Linnaeus, 1767) (Catocalinae)Euclidia mi (Clerck, 1759) (Catocalinae)Lygephila pastinum(Treitschke, 1826) (Catocalinae)Oxyodes scrobiculata(Fabricius, 1775) (Catocalinae)Colocasia coryli(Linnaeus, 1758) (Pantheinae)Diloba caeruleocephala(Linnaeus, 1758) (Pantheinae)Euromoia subpulchra(Alpheraky, 1897) (Pantheinae)Moma alpium(Osbeck, 1778) (Pantheinae)Panthea coenobita(Esper, 1785) (Pantheinae)Raphia frater(Grote, 1864) (Pantheinae)Camptoloma interiorata(Walker, 1865) (Camptolominae)Maurilia iconica (Walker, [1858]) (Chloephorinae)Meganola albula([Denis & Schiffermuller], 1775) (Nolinae)Protodeltote pygarga(Hufnagel, 1766) (Eustrotiinae)Xanthodes albago(Fabricius, 1794) (Bagisarinae)



Table 1 Continued.

Abrostola tripartita (Hufnagel, 1766) [triplasiasensuauct.] (Plusiinae)Autographa gamma(Linnaeus, 1758) (Plusiinae)Acontia lucida(Hufnagel, 1766) (Acontiinae)Aedia leucomelas(Linnaeus, 1758) (Aediinae)Tyta luctuosa([Denis & Schiffermuller], 1775) (Tytinae)Aegle koekeritziana(Hubner, 1799) (Stiriinae)Acronicta rumicis(Linnaeus, 1758) (Acronictinae)Cryphia algae(Fabricius, 1775) (Bryophilinae)Sarbanissa sp.(Agaristinae)Helicoverpa armigera(Hubner, [1809]) (Heliothinae)Heliothis virescens(Fabricius, 1777) (Heliothinae)Pyrrhia umbra(Hufnagel, 1766) (Heliothinae)Agrotis trux (Hubner, [1824]) (Noctuinae)Noctua janthina([Denis & Schiffermuller], 1775) (Noctuinae)Ochropleura plecta(Linnaeus, 1761) (Noctuinae)Peridroma saucia(Hubner, [1808]) (Noctuinae)Xestia xanthographa([Denis & Schiffermuller], 1775) (Noctuinae)Shargacucullia scrophulariae([Denis & Schiffermuller], 1775) (Cuculliinae)Discestra trifolii (Hufnagel, 1766) (Hadeninae)Amphipyra pyramidea(Linnaeus, 1758) (Amphipyrinae)Omphaloscelis lunosa(Haworth, 1809) (Amphipyrinae)Phlogophora meticulosa(Linnaeus, 1758) (Amphipyrinae)

*Many male genitalia of dried specimens belonging to various species of Noctuoidea have been dissected. These species are not included in thepresent list as the exact muscle situation remains unknown to us.

was usually removed prior to the study of the tympanal region In some acronictine, amphipyrine and hadenine species theepaulette is a bizarre structure with acute projections (Fig. 13).(e.g. in Figs 4–6, 8, 9).This type of ctenoid epaulette has been assumed to serve as aphysical barrier to prevent access of the moth ear

3 The membraneous conjunctiva miteMyrmonyssus phalaenodectesTreat, 1954 (Treat, 1957).A structure similar to the true epaulette has been found

The membraneous conjunctiva (Fig. 2, co) forms the enlarged in the catocaline noctuidOxyodes scrobiculataand in theposterior continuation of the metathoracic pleural membrane. chloephorineBena bicolorana. However, in contrast to theNormally, this membrane is not involved in sound perception, epaulette of higher Noctuidae the surface of this pseudepaulettebut in some species it has been modified into an accessoryis smooth.tympanal membrane, e.g. in the plusiineAutographa gamma Still another modification resembling the epaulette was(Eggers, 1919). observed in some Pantheinae. A narrow strip of the conjunctiva

in front of the tympanum is irregularly folded, but notdistinctively elevated.

4 The tympanal bar

The tympanal bar separates the conjunctiva from the adjacent5 The tympanumtympanum. The structure of the tympanal bar is subject to

modifications. Being entirely absent only in the Notodontidaeand the Doidae (Richards, 1932), it takes the form of an The smooth transparent tympanum (5 tympanal membrane)

is usually hidden within the tympanal cavity, but may beelongate shallow groove in the remaining noctuoid families(Fig. 10). Because it is membraneous, the tympanal bar is readily seen in some species. In Notodontidae, the tympanum

is almost always concealed within a deep invagination ofrelatively inconspicuous. Within the Noctuidae, however, it isgenerally sclerotized. The term ‘tympanal sclerite’ is proposed the metepimeron, the epimeral concavity. The notodontid

tympanum occupies a dorsal position within the epimeralfor a sclerotized tympanal bar. A third type of tympanal bar,the epaulette (ep) of Eggers (1919), is clearly restricted to a concavity in a more or less horizontal, roof-like manner.

Internally the distal strand of an auditory chordotonal organsubset of the Noctuidae where the structure has been modifiedinto an elevated ridge subdivided into a series of closely is attached to the delicate tympanal membrane, normally in its

centre (Eggers, 1919; Ghiradella, 1971). Around the point ofadjoining individual lobes. The number and form of theseirregularly folded lobes are subject to considerable interspecific attachment of the chordotonal strand a milky white halo can

be seen within the tympanal membrane.variation (e.g. compare Figs 11 and 12). An SEM of a typicalnoctuid epaulette is shown by Ghiradella (1971). More detailed examination of the tympanal region is possible



Fig. 1. Cladogram showing proposed phylogenetic relationships. For each taxon the approximate number of species has been added in parentheses.

after careful removal of the abdomen. The thorax can then be elaborated dorsally to form a small cavity above the thoraco-abdominal opening. The anterior wall of this cavity is tightlyobserved in posterior view (Fig. 14; see also Fig. 2 in Eggers,

1943). The conjunctiva, the tympanal sclerite (ts) and the fused with the hind wall of the metathoracic scutellum formost of its length. The posterior boundary of the tergal cavitytympanum (ty) are recognizable. In addition, a collar-like

posterior prolongation (cp) that curves around the oval-shaped is demarcated by a thin tergal phragma (tp). This somewhatcomplex morphology may best be observed in a medio-thoraco-abdominal opening (tao) becomes visible (please refer

additionally to Fig. 20 for orientation). The prolongation is sagittal section through the thoraco-abdominal region (Fig. 3).



Fig. 2. Drawing of the tympanal region ofAgrotis exclamationiz(Linnaeus) (Noctuidae, Noctuinae) in lateral view. Abbreviations used: al: alula;co: conjunctiva; cx: eucoxa; e (a, b): metepimeron (a, b); ep: epaulette; es: metepisternum; ls: lateral sclerotization of pleura A 1; m: metameron;poh: postspiracular hood; s 11 2: abdominal sternum 11 2; sc (II, III): scutum (mesonotal, metanotal); sl (II, III): scutellum (mesonotal, metanotal);sp. 1, 2: abdominal spiracles 1, 2; su: subalare; t 1, 2: abdominal terga 1, 2; tr: trochanter; wb: hindwing base.

Table 2. List of synapomorphies. of these, commonly known as the counter-tympanal cavities, aresubject to considerable variation. In some noctuid subfamilies

Clade number Corresponding autapomorphies (no.) (Herminiinae, Rivulinae, Hypenodinae, Hypeninae, Euteliinae,Stictopterinae, Catocalinae, Pantheinae, Camptolominae,

1 Bulbus ejaculatorius elongate (1)Chloephorinae, Sarrothripinae, Nolinae) they are relatively2 Uncus elongate (2)small and are similar to those in the outgroup families (Figs 14,3 Tibial spurs elongate (3a)16). In the remaining noctuid subfamilies they meet mediallyVentral cervical gland present (larva) (3b)

4 Postspiracular hood present (4) (Figs 17–20). Here, the counter-tympanal cavities are5 Male genitalic muscle m.4 split (5) moderately sized (Bagisarinae, Plusiinae, Acontiinae, Stiriinae,6 Counter–tympanal cavities fused medially (6) Sinocharinae, Cuculliinae) (Fig. 17), or are extraordinarily7 SV 2 setae on A 1 absent (larva) (7) large, thereby partly obliterating the thoraco-abdominal opening8 Tympanal bar of epaulette type (8)

(Eustrotiinae, Aediinae, Tytinae, Acronictinae, Bryophilinae,9 A 2 brush organ present (9)

Lophonyctinae, Pseudeustrotiinae, Agaristinae, Heliothinae,10 Corona present on valva (10)Noctuinae, Hadeninae, Amphipyrinae) (Figs 18–20).

Another pair of smooth membranes, the counter-tympana(5 counter-tympanal membranes), are situated medio-dorsallyHowever, the phragma remains incomplete leaving an (oftento the tympanum proper in the metathoracic wall. However,ovoid) central opening to the tergal cavity (tpo). The lateralthe counter-tympanum (ct) normally remains almost entirelywalls of the tergal cavity are invaginated medially to form a

pair of deep cavities. Within the Noctuidae, the size and shape concealed behind the counter-tympanal cavities (Fig. 14).



Fig. 3. Diagrammatic representation of a medio-sagittal section through the thoraco-abdominal region of an adult male ofCatocala fraxini(Linnaeus). The various phragmata and the tergal cavity of the first abdominal segment are indicated. tp: tergal phragma of T 1; c (II, III): coxa II,III; tc: tergal cavity; mmp: meso-metanotal phragma; mp: metanotal phragma; tao: thoraco-abdominal opening; tg: tergal ground. Furtherabbreviations are given in Fig. 2.

The distribution of postspiracular hood and epaulette in the of the Camptolominae–Chloephorinae–Sarrothripinae–Nolinaesubfamily group. A convergent reduction has probably occurredexamined species is given in Table 3.in at least some heliothine species. For example, inHeliothisUnfortunately, different terms have been used by variousvirescensno trace of a postspiracular hood has been found.authors for corresponding tympanal structures. Table 4 lists

the synonymic terms of various authors, and may be usefulAutapomorphy 6. The counter-tympanal cavities of thefor reference to the original papers on tympanal morphology.

outgroup families and of the basal noctuid subfamilies areusually small to medium-sized, and widely separated from

Phylogenetic considerations each other. In contrast, there is a general tendency towardsenlargement of these structures within the higher Noctuidae.

Comparative morphological study of the noctuoid tympanal However, differences in size proved to be continuous ratherregion revealed at least three potentially useful autapomorphiesthan discrete. Consequently, we face difficulties in assigning afor reconstruction of phylogenetic relationships within the clear-cut autapomorphy.Noctuidae. Minet (1986) assumed the enlarged counter-tympanal cavity

an autapomorphy of the Noctuidae1 Aganaidae. However, weAutapomorphy 4. The presence of a prespiracular hood may felt that the most appropriate solution was to recognize a

be considered to represent the ancestral condition within the division above the camptolomine–chloephorine–sarrothripine–Noctuoidea. Lack of a prespiracular hood in the Doidae and noline clade (Fig. 1). Beyond this clade, the counter-tympanalNotodontidae is possibly due to loss (see Miller, 1991; for cavities are not only larger, but are also intimately fuseddetailed discussion). Within the Noctuidae the prespiracular medially. Rather small counter-tympanal cavities which arehood is found only in the Herminiinae. Consequently, the fused medially exist in the subfamilies Bagisarinae, Plusiinae,presence of a postspiracular hood is considered to represent aAcontiinae, Stiriinae, Sinocharinae and Cuculliinae.convincing autapomorphy for the Noctuidae excluding the Enlargement of the counter-tympanal cavities evolvedHerminiinae (Kitching, 1984). The postspiracular hood is independently in part of the Catocalinae (e.g. inOxyodesretained in most noctuid subfamilies and genera. However, it scrobiculata) and also within the Scranciini of the notodontid

subfamily Dudusinae (Miller, 1991).has been reduced to a dorsal remnant in almost all members



Figs 4–9. Noctuoid tympanal regions in lateral aspects (SEM). Orientations are as in Figs 2 & 3. Fig. 4.Lymantria dispar(Linnaeus), Lymantriidae.Fig. 5. Nodaria tristis (Butler), Noctuidae, Herminiinae. Fig. 6.Anoratha costalisMoore, Noctuidae, Hypeninae. Fig. 7.Diloba caeruleocephala(Linnaeus), Noctuidae, Pantheinae. Fig. 8.Meganola strigula([Denis & Schiffermuller)], Noctuidae, Nolinae. Fig. 9.Pheosia gnoma(Fabricius),Notodontidae.

Autapomorphy 8. The tympanal bar is a shallow considered an apomorphy uniting the subfamilies in clade8 (Fig. 1).membraneous groove in the plesiomorphic state. Varying

degrees of sclerotization, sometimes quite indistinct, can be Usually it is quite easy to decide whether an epaulette ispresent or not, but doubts concerning the homology of tympanalobserved around the tympanal groove in basal lineages of the

Noctuidae. The epaulette type of tympanal bar may be bar modifications may arise. For example the epaulette of the



Figs 10–13. Tympanal bars in the Noctuidae (SEM). Fig. 10. Tympanal sclerite ofPenicillaria simplex(Walker), Euteliinae. Fig. 11. Epauletteof Mesoligia furuncula([Denis & Schiffermuller)], Amphipyrinae. Fig. 12. Epaulette ofHeliothis virescens(Fabricius), Heliothinae. Fig. 13.Specialized epaulette ofNacna malachitis(Oberthur), Acronictinae.

cuculliine Shargacucullia scrophulariaeis extremely low and diagrammatically in Fig. 21. The genital segment (i.e.abdominal segment IX) takes the form of a narrow ring thatnot at all ridge-like (Eggers, 1919), but the corrugated surfaceis divided into a dorsal and a ventral semicircle. The dorsalstructure bears resemblance to the typical epaulette. The ctenoidsclerite is termed the tegumen, the ventral one the vinculummodification of the tympanal bar present in a few members of(Klots, 1970). Medioventrally the vinculum is prolonged intothe Acronictinae, Hadeninae and Amphipyrinae is still anothera hollow apodeme of variable size, the saccus. The movableexample of a derived type of epaulette (Treat, 1957).valvae, a large pair of hollow appendages, are attached to theAs already noted above, a smooth pseudepaulette thatlateral margins of the vinculum. The supposed remnant ofprobably evolved independently from the higher noctuidtergum X, the uncus, is dorso-posteriorly connected with theepaulette is present in the catocaline noctuidOxyodestegumen. The tube-like phallus protrudes backwards betweenscrobiculata. Most interestingly, in this same species athe arms of the vinculum. Normally, the phallus is stronglycorresponding enlargement of the counter-tympanal cavities,sclerotized for most of its length. Dorso-anteriorly, the phallusagain probably representing independent evolution, hasbears a membraneous appendage, the bulbus ejaculatoriusoccurred. The correlated occurrence of these structures (i.e.(Fig. 22). The bulbus ejaculatorius has been describedepaulette and enlarged counter-tympanal cavities), observedrepeatedly (e.g. Oiticica, 1946; Callahan, 1960; Mitter, 1988),within the Noctuidae, might indicate functional inter-but has not been widely used as a morphological character. Inrelationship.fact it is almost universally mistaken for the ductusejaculatorius. The bulbus, horseshoe-shaped in cross-section,Character system 2: the male genitaliais attached to the phallus either anteriorly, antero-dorsally, ordorsally. It consists of two intimately approximated contiguousGeneral descriptionlayers (Oiticica, 1946). Apically, the bulbus regularly forms

The structure of the male genitalia and associated musclessome kind of a cap or hood. The structure of the bulbusejaculatorius is subject to the same degree of variation as any(m.1–8) of the superfamily Noctuoidea are depicted


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